Drivers of dune formation control ecosystem function and response to disturbance in a barrier island system.

Barrier islands are landscape features that protect coastlines by reducing wave energy and erosion. Quantifying vegetation-topographic interactions between adjacent habitats are essential for predicting long-term island response and resilience to sea-level rise and disturbance. To understand the effects of dune dynamics on adjacent interior island ecosystem processes, we quantified how sediment availability and previous disturbance regime interact with vegetation to influence dune building and ease of seawater and sediment movement into the island interior on two US mid-Atlantic coast barrier islands. We conducted field surveys of sediment accretion, vegetative cover, and soil characteristics in dune and swale habitats. Digital elevation models provided assessment of water flow resistance from the mean high water mark into the island interior. We found that geographic location impacted sediment accretion rates and Panicum amarum (a species increasing in abundance over time in the Virginia barrier islands) accreted sediment at a significantly lower rate compared to other dune grasses. Dune elevation impacted the ease of seawater flow into the island interior, altering soil chlorides, annual net primary productivity, and soil carbon and nitrogen. Our work demonstrates the importance of incorporating biological processes and cross-island connectivity into future scenario modeling and predictions of rising sea-levels and increased disturbance.

The effects of dune plant roots on loggerhead turtle (Caretta caretta) nest success.

Sand dunes are supported by the extensive root systems of dune plants that anchor the dune and protect it from erosion. While all plants that grow on the dunes support their structure, invasive plants can outcompete the native and non-native dune plants for resources such as nutrients, sunlight, and space to grow. During the summer, sea turtles lay nests on beaches and near dunes; however, their eggs and hatchlings are at risk of destruction and entrapment by dune plant root penetration. Dune plant roots can penetrate sea turtle nest cavities, thus decreasing hatch success of the eggs and emergence success of the hatchlings. We aimed to determine how plant roots impact threatened loggerhead sea turtle (Caretta caretta) nests on Casey Key, Sarasota County, Florida, USA and to assess the factors affecting plant root invasion. Specifically, we determined the effect of plant roots on loggerhead sea turtle nest success, the extent of the impact of invasive plants over non-invasive plants on nests, and if the distance from the dune (barrier) affects whether roots will penetrate the nest. From July to August 2022, we excavated 93 nests to determine the extent of root penetration and identify associated plant species. This field campaign was supported by a long-term dataset (1987-2022) on loggerhead sea turtle nesting across the region. We found that root presence decreased hatch success by 21% and emergence success by 18%, compared to nests that lacked roots within the nest chamber. Nests closer to the dune were more likely to have a higher proportion of root damage and lower hatch and emergence success. This study helps advance understanding of how native and non-native plants affect sea turtle reproductive success and helps inform coastal management aimed at conserving threatened loggerhead populations.

A marine heatwave changes the stabilizing effects of biodiversity in kelp forests.

Biodiversity can stabilize ecological communities through biological insurance, but climate and other environmental changes may disrupt this process via simultaneous ecosystem destabilization and biodiversity loss. While changes to diversity-stability relationships (DSRs) and the underlying mechanisms have been extensively explored in terrestrial plant communities, this topic remains largely unexplored in benthic marine ecosystems that comprise diverse assemblages of producers and consumers. By analyzing two decades of kelp forest biodiversity survey data, we discovered changes in diversity, stability, and their relationships at multiple scales (biological organizational levels, spatial scales, and functional groups) that were linked with the most severe marine heatwave ever documented in the North Pacific Ocean. Moreover, changes in the strength of DSRs during/after the heatwave were more apparent among functional groups than both biological organizational levels (population vs. ecosystem levels) and spatial scales (local vs. broad scales). Specifically, the strength of DSRs decreased for fishes, increased for mobile invertebrates and understory algae, and were unchanged for sessile invertebrates during/after the heatwave. Our findings suggest that biodiversity plays a key role in stabilizing marine ecosystems, but the resilience of DSRs to adverse climate impacts primarily depends on the functional identities of ecological communities.

Dispersal synchronizes giant kelp forests.

Spatial synchrony is the tendency for population fluctuations to be correlated among different locations. This phenomenon is a ubiquitous feature of population dynamics and is important for ecosystem stability, but several aspects of synchrony remain unresolved. In particular, the extent to which any particular mechanism, such as dispersal, contributes to observed synchrony in natural populations has been difficult to determine. To address this gap, we leveraged recent methodological improvements to determine how dispersal structures synchrony in giant kelp (Macrocystis pyrifera), a global marine foundation species that has served as a useful system for understanding synchrony. We quantified population synchrony and fecundity with satellite imagery across 11 years and 880 km of coastline in southern California, USA, and estimated propagule dispersal probabilities using a high-resolution ocean circulation model. Using matrix regression models that control for the influence of geographic distance, resources (seawater nitrate), and disturbance (destructive waves), we discovered that dispersal was an important driver of synchrony. Our findings were robust to assumptions about propagule mortality during dispersal and consistent between two metrics of dispersal: (1) the individual probability of dispersal and (2) estimates of demographic connectivity that incorporate fecundity (the number of propagules dispersing). We also found that dispersal and environmental conditions resulted in geographic clusters with distinct patterns of synchrony. This study is among the few to statistically associate synchrony with dispersal in a natural population and the first to do so in a marine organism. The synchronizing effects of dispersal and environmental conditions on foundation species, such as giant kelp, likely have cascading effects on the spatial stability of biodiversity and ecosystem function.

Spatial synchrony cascades across ecosystem boundaries and up food webs via resource subsidies.

Cross-ecosystem subsidies are critical to ecosystem structure and function, especially in recipient ecosystems where they are the primary source of organic matter to the food web. Subsidies are indicative of processes connecting ecosystems and can couple ecological dynamics across system boundaries. However, the degree to which such flows can induce cross-ecosystem cascades of spatial synchrony, the tendency for system fluctuations to be correlated across locations, is not well understood. Synchrony has destabilizing effects on ecosystems, adding to the importance of understanding spatiotemporal patterns of synchrony transmission. In order to understand whether and how spatial synchrony cascades across the marine-terrestrial boundary via resource subsidies, we studied the relationship between giant kelp forests on rocky nearshore reefs and sandy beach ecosystems that receive resource subsidies in the form of kelp wrack (detritus). We found that synchrony cascades from rocky reefs to sandy beaches, with spatiotemporal patterns mediated by fluctuations in live kelp biomass, wave action, and beach width. Moreover, wrack deposition synchronized local abundances of shorebirds that move among beaches seeking to forage on wrack-associated invertebrates, demonstrating that synchrony due to subsidies propagates across trophic levels in the recipient ecosystem. Synchronizing resource subsidies likely play an underappreciated role in the spatiotemporal structure, functioning, and stability of ecosystems.

Diversity-stability relationships across organism groups and ecosystem types become decoupled across spatial scales.

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity-stability relationship [DSR]). At broader spatial extents, regional-scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long-term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ-diversity on metacommunity variability, but ß-diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ-diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial ß-diversity to reduce spatial synchrony.

Effects of seagrass restoration on coastal fish abundance and diversity.

Restoration is accelerating to reverse global declines of key habitats and recover lost ecosystem functions, particularly in coastal ecosystems. However, there is high uncertainty about the long-term capacity of restored ecosystems to provide habitat and increase biodiversity and the degree to which these ecosystem services are mediated by spatial and temporal environmental variability. We addressed these gaps by sampling fishes biannually for 5-7 years (2012-2018) at 16 sites inside and outside a rapidly expanding restored seagrass meadow in coastal Virginia (USA). Despite substantial among-year variation in abundance and species composition, seine catches in restored seagrass beds were consistently larger (6.4 times more fish, p < 0.001) and more speciose (2.6 times greater species richness, p < 0.001; 3.1 times greater Hill-Shannon diversity, p = 0.03) than seine catches in adjacent unvegetated areas. Catches were particularly larger during summer than autumn (p < 0.01). Structural equation modeling revealed that depth and water residence time interacted to control seagrass presence, leading to higher fish abundance and richness in shallow, well-flushed areas that supported seagrass. Together, our results indicate that seagrass restoration yields large and consistent benefits for many coastal fishes, but that restoration and its benefits are sensitive to the dynamic seascapes in which restoration is conducted. Consideration of how seascape-scale environmental variability affects the success of habitat restoration and subsequent ecosystem function will improve restoration outcomes and the provisioning of ecosystem services.

Efectos de la restauración de pastos marinos sobre la abundancia y diversidad de peces costeros Resumen La restauración ecológica está acelerándose para revertir la declinación mundial de hábitats importantes y para recuperar las funciones ambientales perdidas, particularmente en los ecosistemas costeros. Sin embargo, hay una gran incertidumbre en cuanto a la capacidad a largo plazo que tienen los ecosistemas restaurados de proporcionar hábitats e incrementar la biodiversidad y el grado al que estos servicios ambientales están mediados por la variabilidad ambiental espacial y temporal. Abordamos estos vacíos mediante el muestreo bianual de peces durante 5-7 años (2012-2018) en 16 sitios dentro y fuera de una pradera restaurada de pastos marinos con expansión acelerada en la costa de Virginia (E.U.A.). A pesar de la variación sustancial anual en abundancia y composición de especies, la captura de cerco en los lechos de pastos marinos restaurados fue mayor (6.4 veces más peces, p< 0.001) y con más especies (2.6 veces mayor riqueza de especies, p< 0.001; 3.1 veces mayor diversidad Hill-Shannon, p= 0.03) que la captura de cerco en las áreas aledañas sin vegetación. En particular, las capturas fueron mayores durante el verano que durante el otoño (p < 0.01). Los modelos de ecuaciones estructurales revelaron que la profundidad y el tiempo de residencia acuática interactúan para controlar la presencia de los pastos marinos, lo que resulta en una mayor abundancia y riqueza de peces en áreas someras con buena circulación que fomentan los pastos marinos. En conjunto, nuestros resultados indican que la restauración de los pastos marinos produce grandes beneficios constantes para muchos peces costeros, pero también que la restauración y sus beneficios son sensibles a la dinámica marina en la que se realiza la restauración. Si se considera cómo la variabilidad ambiental a escala de paisaje afecta el éxito de la restauración del hábitat y la función ambiental subsecuente, entonces mejorarán los resultados de restauración y el suministro de servicios ambientales.

Meta-analysis reveals drivers of restoration success for oysters and reef community.

Restoration aims to reverse the global declines of foundation species, but it is unclear how project attributes, the physical setting, and antecedent conditions affect restoration success. In coastal seas worldwide, oyster reef restoration is increasing to counter historical habitat destruction and associated declines in fisheries production and biodiversity. Yet, restoration outcomes are highly variable and the factors that enhance oyster production and nekton abundance and diversity on restored reefs are unresolved. To quantify the drivers of oyster restoration success, we used meta-analysis to synthesize data from 158 restored reefs paired with unstructured habitats along the United States Gulf and Atlantic coasts. The average recovery of oyster production was 65% greater in subtidal (vs. intertidal) zones, 173% greater in polyhaline (vs. mesohaline) environments and increased with tidal range, demonstrating that physical conditions can strongly influence the restoration success of foundation species. Additionally, restoration increased the relative abundance and richness of nektonic fishes and invertebrates over time as reefs aged (at least 8 years post-construction). Thus, the restoration benefits for provisioning habitat and enhancing biodiversity accrue over time, highlighting that restoration projects need multiple years to maximize ecosystem functions. Furthermore, long-term monitoring of restored and control sites is needed to assess restoration outcomes and associated drivers. Last, our work reveals data constraints for several potential drivers of restoration outcomes, including reef construction material, reef dimensions, harvest pressure and disease prevalence. More experimental and observational studies are needed to target these factors and measure them with consistent methods across studies. Our findings indicate that the assisted recovery of foundation species yields several enhancements to ecosystem services, but such benefits are mediated by time and environmental conditions.

Meta-analysis of ecosystem services associated with oyster restoration.

Restoration of foundation species promises to reverse environmental degradation and return lost ecosystem services, but a lack of standardized evaluation across projects limits understanding of recovery, especially in marine systems. Oyster reefs are restored to reverse massive global declines and reclaim valuable ecosystem services, but the success of these projects has not been systematically and comprehensively quantified. We synthesized data on ecosystem services associated with oyster restoration from 245 pairs of restored and degraded reefs and 136 pairs of restored and reference reefs across 3500 km of U.S. Gulf of Mexico and Atlantic coastlines. On average, restoration was associated with a 21-fold increase in oyster production (mean log response ratio = 3.08 [95% confidence interval: 2.58-3.58]), 34-97% enhancement of habitat provisioning (mean community abundance = 0.51 [0.41-0.61], mean richness = 0.29 [0.19-0.39], and mean biomass = 0.69 [0.39-0.99]), 54% more nitrogen removal (mean = 0.43 [0.13-0.73]), and 89-95% greater sediment nutrients (mean = 0.67 [0.27-1.07]) and organic matter (mean = 0.64 [0.44-0.84]) relative to degraded habitats. Moreover, restored reefs matched reference reefs for these ecosystem services. Our results support the continued and expanded use of oyster restoration to enhance ecosystem services of degraded coastal systems and match many functions provided by reference reefs.

La restauración de especies fundadoras promete revertir la degradación ambiental y restituir servicios ambientales perdidos, pero la falta de evaluación estandarizada de proyectos limita la comprensión de la recuperación, especialmente en sistemas marinos. Los bancos de ostión son restaurados para revertir declinaciones globales masivas y recuperar servicios ecosistémicos valiosos, pero el éxito de estos proyectos no ha sido cuantificado sistemática ni integralmente. Sintetizamos datos sobre los servicios ecosistémicos asociados con la restauración de ostiones de 245 pares de bancos restaurados y degradados y 136 pares de bancos restaurados y de referencia a lo largo de 3500 km de costa del Golfo de México y Atlántico norteamericanos. En promedio, la restauración se asoció con un incremento de 21 veces en la producción de ostión (media de proporción de respuesta log = 3.08 [95% IC 2.58-3.58]), mejoras entre 34 y 97% en el aprovisionamiento de hábitat (abundancia media = 0.51 [0.41-0.61], riqueza media = 0.29 [0.19-0.39], y biomasa media = 0.69 [0.39-0.99]), 54% más remoción de nitrógeno (media = 0.43 [0.13-0.73]), y 89-95% más nutrientes en sedimento (media = 0.67 [0.27-1.07]) y materia orgánica (media = 0.64 [0.44-0.84]) en relación con hábitats degradados. Más aun, estos servicios ecosistémicos de los bancos restaurados fueron muy similares en los bancos de referencia. Nuestros resultados sustentan el uso continuo y expandido de la restauración de ostiones para mejorar los servicios ecosistémicos de sistemas costeros degradados y que sean parecidos a las numerosas funciones proporcionadas por los bancos de referencia.

Disturbance and nutrients synchronise kelp forests across scales through interacting Moran effects.

Spatial synchrony is a ubiquitous and important feature of population dynamics, but many aspects of this phenomenon are not well understood. In particular, it is largely unknown how multiple environmental drivers interact to determine synchrony via Moran effects, and how these impacts vary across spatial and temporal scales. Using new wavelet statistical techniques, we characterised synchrony in populations of giant kelp Macrocystis pyrifera, a widely distributed marine foundation species, and related synchrony to variation in oceanographic conditions across 33 years (1987-2019) and >900 km of coastline in California, USA. We discovered that disturbance (storm-driven waves) and resources (seawater nutrients)-underpinned by climatic variability-act individually and interactively to produce synchrony in giant kelp across geography and timescales. Our findings demonstrate that understanding and predicting synchrony, and thus the regional stability of populations, relies on resolving the synergistic and antagonistic Moran effects of multiple environmental drivers acting on different timescales.

Tail-dependent spatial synchrony arises from nonlinear driver-response relationships.

Spatial synchrony may be tail-dependent, that is, stronger when populations are abundant than scarce, or vice-versa. Here, 'tail-dependent' follows from distributions having a lower tail consisting of relatively low values and an upper tail of relatively high values. We present a general theory of how the distribution and correlation structure of an environmental driver translates into tail-dependent spatial synchrony through a non-linear response, and examine empirical evidence for theoretical predictions in giant kelp along the California coastline. In sheltered areas, kelp declines synchronously (lower-tail dependence) when waves are relatively intense, because waves below a certain height do little damage to kelp. Conversely, in exposed areas, kelp is synchronised primarily by periods of calmness that cause shared recovery (upper-tail dependence). We find evidence for geographies of tail dependence in synchrony, which helps structure regional population resilience: areas where population declines are asynchronous may be more resilient to disturbance because remnant populations facilitate reestablishment.

The long and the short of it: Mechanisms of synchronous and compensatory dynamics across temporal scales.

Synchronous dynamics (fluctuations that occur in unison) are universal phenomena with widespread implications for ecological stability. Synchronous dynamics can amplify the destabilizing effect of environmental variability on ecosystem functions such as productivity, whereas the inverse, compensatory dynamics, can stabilize function. Here we combine simulation and empirical analyses to elucidate mechanisms that underlie patterns of synchronous versus compensatory dynamics. In both simulated and empirical communities, we show that synchronous and compensatory dynamics are not mutually exclusive but instead can vary by timescale. Our simulations identify multiple mechanisms that can generate timescale-specific patterns, including different environmental drivers, diverse life histories, dispersal, and non-stationary dynamics. We find that traditional metrics for quantifying synchronous dynamics are often biased toward long-term drivers and may miss the importance of short-term drivers. Our findings indicate key mechanisms to consider when assessing synchronous versus compensatory dynamics and our approach provides a pathway for disentangling these dynamics in natural systems.

Disturbance structures canopy and understory productivity along an environmental gradient.

Disturbances often disproportionately impact different vegetation layers in forests and other vertically stratified ecosystems, shaping community structure and ecosystem function. However, disturbance-driven changes may be mediated by environmental conditions that affect habitat quality and species interactions. In a decade-long field experiment, we tested how kelp forest net primary productivity (NPP) responds to repeated canopy loss along a gradient in grazing and substrate suitability. We discovered that habitat quality can mediate the effects of intensified disturbance on canopy and understory NPP. Experimental annual and quarterly disturbances suppressed total macroalgal NPP, but effects were strongest in high-quality habitats that supported dense kelp canopies that were removed by disturbance. Understory macroalgae partly compensated for canopy NPP losses and this effect magnified with increasing habitat quality. Disturbance-driven increases in understory NPP were still rising after 5-10 years of disturbance, demonstrating the value of long-term experimentation for understanding ecosystem responses to changing disturbance regimes.

The spatial synchrony of species richness and its relationship to ecosystem stability.

Synchrony is broadly important to population and community dynamics due to its ubiquity and implications for extinction dynamics, system stability, and species diversity. Investigations of synchrony in community ecology have tended to focus on covariance in the abundances of multiple species in a single location. Yet, the importance of regional environmental variation and spatial processes in community dynamics suggests that community properties, such as species richness, could fluctuate synchronously across patches in a metacommunity, in an analog of population spatial synchrony. Here, we test the prevalence of this phenomenon and the conditions under which it may occur using theoretical simulations and empirical data from 20 marine and terrestrial metacommunities. Additionally, given the importance of biodiversity for stability of ecosystem function, we posit that spatial synchrony in species richness is strongly related to stability. Our findings show that metacommunities often exhibit spatial synchrony in species richness. We also found that richness synchrony can be driven by environmental stochasticity and dispersal, two mechanisms of population spatial synchrony. Richness synchrony also depended on community structure, including species evenness and beta diversity. Strikingly, ecosystem stability was more strongly related to richness synchrony than to species richness itself, likely because richness synchrony integrates information about community processes and environmental forcing. Our study highlights a new approach for studying spatiotemporal community dynamics and emphasizes the spatial dimensions of community dynamics and stability.

Disturbance size and frequency mediate the coexistence of benthic spatial competitors.

Disturbance plays a key role in structuring community dynamics and is central to conservation and natural resource management. However, ecologists continue to debate the importance of disturbance for species coexistence and biodiversity. Such disagreements may arise in part because few studies have examined variation across multiple dimensions of disturbance (e.g., size, frequency) and how the effects of disturbance may depend on species attributes (e.g., competitiveness, dispersal ability). In light of this gap in understanding and accelerating changes to disturbance regimes worldwide, we used spatial population models to explore how disturbance size and frequency interact with species attributes to affect coexistence between seagrass (Zostera marina) and colonial burrowing shrimp (Neotrypaea californiensis) that compete for benthic space in estuaries throughout the west coast of North America. By simulating population dynamics under a range of ecologically relevant disturbance regimes, we discovered that intermediate disturbance (approximately 9-23% of landscape area per year) to short-dispersing, competitively dominant seagrass can foster long-term stable coexistence with broad-dispersing, competitively inferior burrowing shrimp via the spatial storage effect. When holding the total extent of disturbance constant, the individual size and annual frequency of disturbance altered landscape spatial patterns and mediated the dominance and evenness of competitors. Many small disturbances favored short-dispersing seagrass by hastening recolonization, whereas fewer large disturbances benefited rapidly colonizing burrowing shrimp by creating temporary refugia from competition. As a result, large, infrequent disturbances generally improved the strength and stability of coexistence relative to small, frequent disturbances. Regardless of disturbance size or frequency, the dispersal ability of the superior competitor (seagrass), the competitive ability of the inferior competitor (burrowing shrimp), and the reproduction and survival of both species strongly influenced population abundances and coexistence. Our results show that disturbance size and frequency can promote or constrain coexistence by altering the duration of time over which inferior competitors can escape competitive exclusion, particularly when colonization depends on the spatial pattern of disturbance due to dispersal traits. For coastal managers and conservation practitioners, our findings indicate that reducing particularly large disturbances may help conserve globally imperiled seagrass meadows and control burrowing shrimp colonies that can threaten the viability of oyster aquaculture.

Loss of foundation species: disturbance frequency outweighs severity in structuring kelp forest communities.

Disturbances often cause the disproportionate loss of foundation species but understanding how the frequency and severity of disturbance to such organisms influence biological communities remains unresolved. This gap in knowledge exists in part because of the rarity of ecologically meaningful studies capable of disentangling different elements of disturbance. Hence, we carried out a long-term (9 yr), large-scale (2,000 m2 plots), spatially replicated (4 sites) field experiment in which we manipulated disturbance to a globally distributed marine foundation species, the giant kelp Macrocystis pyrifera, and tracked community responses over time. To distinguish the effects of disturbance frequency and severity on the biodiversity and composition of temperate rocky reef communities, we simulated the repeated loss of giant kelp from destructive winter waves across a background of natural variation in disturbance. By following the response of over 200 taxa from the surrounding community, we discovered that the frequency of disturbance to giant kelp changed the biomass, diversity, and composition of community guilds in a manner commensurate with their dependence on the physical (i.e., benthic light and space), trophic (i.e., living and detrital biomass), and habitat (i.e., biogenic structure) resources mediated by this foundation species. Annual winter disturbance to giant kelp reduced living and detrital giant kelp biomass by 57% and 40%, respectively, enhanced bottom light by 22%, and halved the seafloor area covered by giant kelp holdfasts. Concomitantly, the biomass of understory algae and epilithic sessile invertebrates more than doubled, while the biomass of rock-boring clams, mobile invertebrates, and fishes decreased 30-61%. Frequent loss of giant kelp boosted understory algal richness by 82% and lowered sessile invertebrate richness by 13% but did not affect the biodiversity of mobile fauna. In contrast to changes driven by disturbance frequency, interannual variation in the severity of disturbance to giant kelp had weaker, less consistent effects, causing only modest changes in assemblages of sessile invertebrates, mobile invertebrate herbivores, and fishes. Our results broaden the foundation species concept by demonstrating that repeated disturbance to a dominant habitat-forming species can outweigh the influence of less frequent but severe disturbances for the surrounding community.

Fluctuations in population fecundity drive variation in demographic connectivity and metapopulation dynamics.

Demographic connectivity is vital to sustaining metapopulations yet often changes dramatically through time due to variation in the production and dispersal of offspring. However, the relative importance of variation in fecundity and dispersal in determining the connectivity and dynamics of metapopulations is poorly understood due to the paucity of comprehensive spatio-temporal data on these processes for most species. We quantified connectivity in metapopulations of a marine foundation species (giant kelp Macrocystis pyrifera) across 11 years and approximately 900 km of coastline by estimating population fecundity with satellite imagery and propagule dispersal using a high-resolution ocean circulation model. By varying the temporal complexity of different connectivity measures and comparing their ability to explain observed extinction-colonization dynamics, we discovered that fluctuations in population fecundity, rather than fluctuations in dispersal, are the dominant driver of variation in connectivity and contribute substantially to metapopulation recovery and persistence. Thus, for species with high variability in reproductive output and modest variability in dispersal (most plants, many animals), connectivity measures ignoring fluctuations in fecundity may overestimate connectivity and likelihoods of persistence, limiting their value for understanding and conserving metapopulations. However, we demonstrate how connectivity measures can be simplified while retaining utility, validating a practical solution for data-limited systems.

Metabolomics Reveals Cryptic Interactive Effects of Species Interactions and Environmental Stress on Nitrogen and Sulfur Metabolism in Seagrass.

Eutrophication of estuaries and coastal seas is accelerating, increasing light stress on subtidal marine plants and changing their interactions with other species. To date, we have limited understanding of how such variations in environmental and biological stress modify the impact of interactions among foundational species and eventually affect ecosystem health. Here, we used metabolomics to assess the impact of light reductions on interactions between the seagrass Zostera marina, an important habitat-forming marine plant, and the abundant and commercially important blue mussel Mytilus edulis. Plant performance varied with light availability but was unaffected by the presence of mussels. Metabolomic analysis, on the other hand, revealed an interaction between light availability and presence of M. edulis on seagrass metabolism. Under high light, mussels stimulated seagrass nitrogen and energy metabolism. Conversely, in low light mussels impeded nitrogen and energy metabolism, and enhanced responses against sulfide toxicity, causing inhibited oxidative energy metabolism and tissue degradation. Metabolomic analysis thereby revealed cryptic changes to seagrass condition that could not be detected by traditional approaches. Our findings suggest that coastal eutrophication and associated reductions in light may shift seagrass-bivalve interactions from mutualistic to antagonistic, which is important for conservation management of seagrass meadows.

Invasive prey indirectly increase predation on their native competitors.

Ecological theory predicts that invasive prey can interact with native prey directly by competing for shared resources or indirectly by changing the abundance or behavior of shared native predators. However, both the study and management of invasive prey have historically overlooked indirect effects. In southern California estuaries, introduction of the Asian nest mussel Arcuatula senhousia has been linked to profound changes in native bivalve assemblages, but the mechanisms of these interactions remain unclear. We performed three field experiments to assess the mechanisms of competition between Arcuatula and native bivalves, and evaluated the potential for Arcuatula to indirectly mediate native predator-prey dynamics. We found that Arcuatula reduces the diversity, abundance, and size of native bivalve recruits by preemptively exploiting space in surface sediments. When paired with native shallow-dwelling clams (Chione undatella and Laevicardium substriatum), Arcuatula reduces adult survival through overgrowth competition. However, Arcuatula also attracts native predators, causing apparent competition by indirectly increasing predation of native clams, especially for poorly defended species. Therefore, invasive prey can indirectly increase predation rates on native competitors by changing the behavior of shared native predators, but the magnitude of apparent competition strongly depends on the vulnerability of natives to predation. Interestingly, our results indicate that the vulnerability of invasive prey to predation can greatly exacerbate impacts on their native competitors. Our findings suggest that consideration of both direct and indirect effects of invasive prey, as well as native predator-prey relationships, should lead to more effective invasive species management.

Connectivity structures local population dynamics: a long-term empirical test in a large metapopulation system.

Ecological theory predicts that demographic connectivity structures the dynamics of local populations within metapopulation systems, but empirical support has been constrained by major limitations in data and methodology. We tested this prediction for giant kelp Macrocystis pyrifera, a key habitat-forming species in temperate coastal ecosystems worldwide, in southern California, USA. We combined a long-term (22 years), large-scale (~500 km coastline), high-resolution census of abundance with novel patch delineation methods and an innovative connectivity measure incorporating oceanographic transport and source fecundity. Connectivity strongly predicted local dynamics (well-connected patches had lower probabilities of extinction and higher probabilities of colonization, leading to greater likelihoods of occupancy) but this relationship was mediated by patch size. Moreover, the relationship between connectivity and local population dynamics varied over time, possibly due to temporal variation in oceanographic transport processes. Surprisingly, connectivity had a smaller influence on colonization relative to extinction, possibly because local ecological factors differ greatly between extinct and extant patches. Our results provide the first comprehensive evidence that southern California giant kelp populations function as a metapopulation system, challenging the view that populations of this important foundation species are governed exclusively by self-replenishment.